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Page last updated
Wed 1 October 2025

The Freshwater Mussels (Unionoida) of the World (and other less consequential bivalves)

FM(U)otW(aolcb) is the web version of the MUSSEL Project Database. Follow the links to browse the data or use the custom Google search field. Either way, you win!

Search the database pages:

Bivalvia | Unionoida | Unionoidea

family MARGARITIFERIDAE Henderson, 1929: cladomics
     6 genera, 2 subfamilies

Cladomics pages are experimental — even more so than the rest of this site!

The purpose of the cladomics pages for each genus and family-group level taxon is to provide statistics and references indicating the extent to which published phylogenetic work supports the classification used here.

The Cladomics Summary provides an overview of the number of studies that have included terminal species representing this taxon as well as an indication of how frequently it has been recovered as monophyletic.

Following the summary, the available published clades representing the taxon are listed.

For more information, visit the Cladomics FARQ (frequently asked rhetorical questions) page. Abbreviations for the various statistics are explained on the FARQ!

Cladomics Summary

number of species = 16 (43 total)
number of subtaxa (rank: subfamily) = 8
analyzed in 259 cladograms in 146 publications
maximum number of terminals in an analysis = 16 (mean = 3.15, median = 2)
maximum number of subtaxa (subfamily) = 2 (mean = 1.26, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 174
maximum number of included terminals = 16, (mean = 4.2 median = 2)
maximum number of subtaxa (subfamily) = 2 (mean = 1.39, median = 1)
proportion of cladograms that support basic monophyly = 0.95

Rigorous Monophyly

number of cladograms that test rigorous monophyly = 65
(Rigorous monophyly refers to cladograms with terminal species representing more than one subtaxon (subfamily) taxon and at least one rigorous outgroup (family).
maximum number of included terminals = 15 (mean = 6.26, median = 6)
maximum number of subtaxa (subfamily) = 2 (mean = 2, median = 2)
proportion of cladograms that support rigorous monophyly = 0.97

Pairwise Cladogram Clustering

The graphs below depict in two dimensions all pair-wise ingroup distances within the cladome. See the Cladmoics FARQ for more information.


PCoA Ordination


NJ Tree

Zhou et al., 2007 [Publ 2658], fig. 1

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
321192181810
  +-------------------- Margaritiferinae **       
  |                                            
  |  +----------------- Anodontini             
--+  |                                         
  |  |     +----------- Chamberlainiini        
  |  |  +--+                                   
  +--+  |  |  +-------- Lamprotula_leaii       
     |  |  +--+                                
     |  |     |  +----- Lamprotula_caveata     
     |  |     +--+                             
     |  |        |  +-- Gibbosulinae **           
     +--+        +--+                          
        |           +-- Lamprotula_cornuumlunae
        |                                      
        |        +----- Aculamprotula          
        |     +--+                             
        |     |  |  +-- Tchangsinaia           
        |     |  +--+                          
        +-----+     +-- Nodularia              
              |                                
              |  +----- Schistodesmus          
              +--+                             
                 |  +-- Lanceolariini          
                 +--+                          
                    +-- Acuticostini

Huang et al., 2019 [Publ 3843], fig. 2

** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
131392186160128
  +-------- Unionidae               
--+                                 
  |  +----- Gibbosula_rochechouartii **
  +--+                              
     |  +-- Gibbosula_laosensis **     
     +--+                           
        +-- Margaritiferinae **

Lyubas et al., 2019 [Publ 3963], fig. 10

iTtTbiriSoTroTroSiD
161612200031
  +-- Gibbosulinae **    
--+                   
  +-- Margaritiferinae **

Chen et al., 2025 [Publ 4455], fig. 1

iTtTbiriSoTroTroSiD
1515132112130
  +-------------- Unionidae                
  |                                        
--+  +----------- Pseudunio_marocanus **      
  |  |                                     
  |  |        +-- Pseudunio_auricularius__1 **
  +--+        |                            
     |  +-----+-- Pseudunio_auricularius__2 **
     |  |     |                            
     +--+     +-- Pseudunio_homsensis **      
        |                                  
        |  +----- Gibbosulinae **             
        +--+                               
           |  +-- Cumberlandia **             
           +--+                            
              +-- Margaritifera **

Lopes-Lima et al., 2018 [Publ 3829], fig. 1

iTtTbiriSoTroTroSiD
1414122143129
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Lopes-Lima et al., 2018 [Publ 3829], fig. 3

iTtTbiriSoTroTroSiD
1414122143129
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Huang et al., 2019 [Publ 3843], fig. S4

iTtTbiriSoTroTroSiD
131392186160128
  +-------- Unionidae               
--+                                 
  |  +----- Gibbosula_rochechouartii **
  +--+                              
     |  +-- Gibbosula_laosensis **     
     +--+                           
        +-- Margaritiferinae **

Huang et al., 2019 [Publ 3843], fig. S3

iTtTbiriSoTroTroSiD
1313112184158128
  +-------- Unionidae               
--+                                 
  |  +----- Gibbosula_rochechouartii **
  +--+                              
     |  +-- Gibbosula_laosensis **     
     +--+                           
        +-- Margaritiferinae **

Sano et al., 2020 [Publ 4138], fig. 4

iTtTbiriSoTroTroSiD
12121025645129
  +----------- Unionidae    
--+                         
  |  +-------- Cumberlandia ** 
  +--+                      
     |  +----- Pseudunio **    
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Margaritifera **

Wu et al., 2020 [Publ 4121], fig. 2

iTtTbiriSoTroTroSiD
12121022211129
  +----------- Unionidae    
--+                         
  |  +-------- Cumberlandia ** 
  +--+                      
     |  +----- Margaritifera **
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Pseudunio **

Sano et al., 2020 [Publ 4138], fig. 2

iTtTbiriSoTroTroSiD
12121025645129
  +----------- Unionidae    
--+                         
  |  +-------- Cumberlandia ** 
  +--+                      
     |  +----- Margaritifera **
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Pseudunio **

Sano et al., 2020 [Publ 4138], fig. 3

iTtTbiriSoTroTroSiD
12121025645129
  +----------- Unionidae    
--+                         
  |  +-------- Cumberlandia ** 
  +--+                      
     |  +----- Pseudunio **    
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Margaritifera **

Huang et al., 2018 [Publ 3828], fig. 3_5

iTtTbiriSoTroTroSiD
1212102176127
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wu et al., 2020 [Publ 4121], fig. S1

iTtTbiriSoTroTroSiD
12121022211127
  +-------- Unionidae               
--+                                 
  |  +----- Gibbosula_rochechouartii **
  +--+                              
     |  +-- Gibbosula_laosensis **     
     +--+                           
        +-- Margaritiferinae **

Araujo et al., 2016 [Publ 3746], fig. 2a

iTtTbiriSoTroTroSiD
11119200029
  +-------- Cumberlandia ** 
--+                      
  |  +----- Pseudunio **    
  +--+                   
     |  +-- Gibbosulinae ** 
     +--+                
        +-- Margaritifera **

Araujo et al., 2016 [Publ 3746], fig. 1_2b

iTtTbiriSoTroTroSiD
11119221127
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Vikhrev et al., 2018 [Publ 3812], fig. 3

iTtTbiriSoTroTroSiD
11119222127
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Bolotov et al., 2016 [Publ 3743], fig. 4

iTtTbiriSoTroTroSiD
101082204128
  +----------- Unionidae    
--+                         
  |  +-------- Pseudunio **    
  +--+                      
     |  +----- Margaritifera **
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Cumberlandia **

Bolotov et al., 2017 [Publ 3761], fig. S2

iTtTbiriSoTroTroSiD
101072129114128
  +-------- Unionidae    
  |                      
--+  +----- Cumberlandia ** 
  |  |                   
  +--+  +-- Gibbosulinae ** 
     |  |                
     +--+-- Pseudunio **    
        |                
        +-- Margaritifera **

Bolotov et al., 2017 [Publ 3761], fig. S1

iTtTbiriSoTroTroSiD
101082129114128
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae ** 
     +--+                
        +-- Pseudunio **

Jeratthitikul et al., 2022 [Publ 4260], fig. S2_2

iTtTbiriSoTroTroSiD
1010826666128
  +----------- Unionidae    
--+                         
  |  +-------- Cumberlandia ** 
  +--+                      
     |  +----- Margaritifera **
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Pseudunio **

Jeratthitikul et al., 2022 [Publ 4260], fig. S1_1

iTtTbiriSoTroTroSiD
1010826666128
  +----------- Unionidae    
--+                         
  |  +-------- Cumberlandia ** 
  +--+                      
     |  +----- Gibbosulinae ** 
     +--+                   
        |  +-- Pseudunio **    
        +--+                
           +-- Margaritifera **

Bolotov et al., 2016 [Publ 3743], fig. 2

iTtTbiriSoTroTroSiD
101082204128
  +----------- Unionidae    
--+                         
  |  +-------- Pseudunio **    
  +--+                      
     |  +----- Margaritifera **
     +--+                   
        |  +-- Gibbosulinae ** 
        +--+                
           +-- Cumberlandia **

Takeuchi et al., 2015 [Publ 3661], fig. 2

iTtTbiriSoTroTroSiD
886121125
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Takeuchi et al., 2015 [Publ 3661], fig. 3

iTtTbiriSoTroTroSiD
886121125
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Bolotov et al., 2015 [Publ 3646], fig. 2B

iTtTbiriSoTroTroSiD
886111125
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Bolotov et al., 2015 [Publ 3646], fig. 2A

iTtTbiriSoTroTroSiD
885111125
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Inoue et al., 2013 [Publ 3579], fig. 3

iTtTbiriSoTroTroSiD
886111125
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Huff et al., 2004 [Publ 2697], fig. 2

iTtTbiriSoTroTroSiD
773153125
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Wu et al., 2021 [Publ 4366], fig. 3-ML

iTtTbiriSoTroTroSiD
77526662122
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Araujo et al., 2009 [Publ 2395], fig. 3

iTtTbiriSoTroTroSiD
775111124
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Wu et al., 2021 [Publ 4366], fig. 4

iTtTbiriSoTroTroSiD
77526157122
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Huff et al., 2004 [Publ 2697], fig. 5-6

iTtTbiriSoTroTroSiD
775153124
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Wu et al., 2021 [Publ 4366], fig. 3-BI

iTtTbiriSoTroTroSiD
77526662122
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wu et al., 2023 [Publ 4339], fig. 4_S5-nt

iTtTbiriSoTroTroSiD
77528884122
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Guerra et al., 2019 [Publ 4168], fig. 1

iTtTbiriSoTroTroSiD
77523421123
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Gomes-dos-Santos et al., 2019 [Publ 3943], fig. 1

iTtTbiriSoTroTroSiD
775233122
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Huff et al., 2004 [Publ 2697], fig. 4

iTtTbiriSoTroTroSiD
774143124
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Wu et al., 2021 [Publ 4366], fig. 2_I

iTtTbiriSoTroTroSiD
77528474123
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Guerra et al., 2019 [Publ 4168], fig. 2

iTtTbiriSoTroTroSiD
77523421123
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wu et al., 2021 [Publ 4366], fig. 6

iTtTbiriSoTroTroSiD
77528474123
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wu et al., 2023 [Publ 4339], fig. 4_S5-aa

iTtTbiriSoTroTroSiD
77528884122
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Dai et al., 2025 [Publ 4440], fig. 3

iTtTbiriSoTroTroSiD
66428279121
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Huff et al., 2004 [Publ 2697], fig. 3

iTtTbiriSoTroTroSiD
662121123
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Walker et al., 2006 [Publ 2666], fig. 6

iTtTbiriSoTroTroSiD
66413010123
  +-------- Unioninae       
--+                         
  |  +----- Margaritiferinae **
  +--+                      
     |  +-- Gonideinae      
     +--+                   
        +-- Ambleminae

Chen et al., 2025 [Publ 4448], fig. 3

iTtTbiriSoTroTroSiD
66427976121
  +-- Gibbosulinae **    
--+                   
  +-- Margaritiferinae **

Wu et al., 2022 [Publ 4238], fig. 3

iTtTbiriSoTroTroSiD
66426764121
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Huang et al., 2018 [Publ 3828], fig. 6

iTtTbiriSoTroTroSiD
66423632121
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Huang et al., 2019 [Publ 3843], fig. 3

iTtTbiriSoTroTroSiD
66425046121
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Bolotov et al., 2020 [Publ 4100], fig. 1

iTtTbiriSoTroTroSiD
5502113113123
  +----- Unionidae                  
  |                                 
  |  +-- Gibbosulinae **               
--+  |                              
  |  +-- Margaritifera_dahurica **     
  |  |                              
  +--+-- Margaritifera_margaritifera **
     |                              
     +-- Margaritifera_laevis **       
     |                              
     +-- Margaritifera_middendorffi **

Bolotov et al., 2020 [Publ 4084], fig. 1

iTtTbiriSoTroTroSiD
44013232123
  +----- Margaritiferinae **
  |                      
--+----- Ambleminae      
  |                      
  |  +-- Gonideinae      
  +--+                   
     +-- Unioninae

Chase et al., 2018 [Publ 3847], fig. 4

iTtTbiriSoTroTroSiD
44213433121
1

Breton et al., 2011 [Publ 3376], fig. 3

iTtTbiriSoTroTroSiD
44213332121
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Wu et al., 2019 [Publ 3864], fig. 3

iTtTbiriSoTroTroSiD
44224240120
  +-- Gibbosulinae **    
--+                   
  +-- Margaritiferinae **

Wu et al., 2020 [Publ 4121], fig. 3

iTtTbiriSoTroTroSiD
44224541120
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Fukata & Iigo, 2020 [Publ 4106], fig. 1

iTtTbiriSoTroTroSiD
33125050121
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Cha et al., 2018 [Publ 3916], fig. 1

iTtTbiriSoTroTroSiD
33113535123
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Rosenberg et al., 1997 [Publ 2720], fig. 2A

iTtTbiriSoTroTroSiD
322411918137
  +----- Cumberlandia ** 
  |                   
  +----- None__2      
--+                   
  +----- Margaritifera **
  |                   
  |  +-- None__1      
  +--+                
     +-- Unionidae

Lopes-Lima et al., 2017 [Publ 3792], fig. 2

iTtTbiriSoTroTroSiD
33112929123
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Aunins et al., 2018 [Publ 3863], fig. 1

iTtTbiriSoTroTroSiD
33113737121
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Machordom et al., 2003 [Publ 2688], fig. 3

iTtTbiriSoTroTroSiD
331111120
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Zieritz et al., 2021 [Publ 4231], fig. 3-F_5

iTtTbiriSoTroTroSiD
33114545123
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lydeard et al., 1996 [Publ 2656], fig. 3

iTtTbiriSoTroTroSiD
33112424120
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Sano et al., 2020 [Publ 4138], fig. 1

iTtTbiriSoTroTroSiD
33123130121
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wu et al., 2020 [Publ 4147], fig. 4

iTtTbiriSoTroTroSiD
33121111120
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Lydeard et al., 1996 [Publ 2656], fig. 4

iTtTbiriSoTroTroSiD
33112524120
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Stewart et al., 2021 [Publ 4451], fig. 1

iTtTbiriSoTroTroSiD
33111313122
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Froufe et al., 2019 [Publ 3936], fig. 3

iTtTbiriSoTroTroSiD
33114141123
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Froufe et al., 2019 [Publ 3936], fig. 6

iTtTbiriSoTroTroSiD
33114141123
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Mock et al., 2013 [Publ 3598], fig. 3

iTtTbiriSoTroTroSiD
331100022
1

Guerra et al., 2019 [Publ 4168], fig. 1-M

iTtTbiriSoTroTroSiD
33111917120
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Rosenberg et al., 1994 [Publ 2719], fig. 3

iTtTbiriSoTroTroSiD
319511616136
  +-- Cumberlandia ** 
  |                
--+-- Margaritifera **
  |                
  +-- Unionidae

Walker et al., 2006 [Publ 2666], fig. 6-M

iTtTbiriSoTroTroSiD
33111110120
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Gomes-dos-Santos et al., 2019 [Publ 3943], fig. 1-M

iTtTbiriSoTroTroSiD
331133120
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Rosenberg et al., 1997 [Publ 2720], fig. 2B

iTtTbiriSoTroTroSiD
322311918136
  +-- Cumberlandia ** 
  |                
  +-- None__1      
  |                
--+-- None__2      
  |                
  +-- Margaritifera **
  |                
  +-- Unionidae

Guerra et al., 2019 [Publ 4168], fig. 2-M

iTtTbiriSoTroTroSiD
33111917120
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Zieritz et al., 2024 [Publ 4381], fig. 2

iTtTbiriSoTroTroSiD
220144441--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lopes-Lima et al., 2020 [Publ 4080], fig. 8

iTtTbiriSoTroTroSiD
2201551--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Huang et al., 2002 [Publ 2657], fig. 2

iTtTbiriSoTroTroSiD
234013232139
  +-- Cumberlandia ** 
  |                
--+-- Margaritifera **
  |                
  +-- Unionidae

Hoeh et al., 2001 [Publ 2652], fig. 14f4

iTtTbiriSoTroTroSiD
220129181--
  +----------- Parreysiinae    
--+                            
  |  +-------- Margaritiferinae **
  +--+                         
     |  +----- Gonideinae      
     +--+                      
        |  +-- Unioninae       
        +--+                   
           +-- Ambleminae

Konopleva et al., 2017 [Publ 3807], fig. 6

iTtTbiriSoTroTroSiD
220129211--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Wu et al., 2024 [Publ 4379], fig. 5

iTtTbiriSoTroTroSiD
220251511--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Sano et al., 2017 [Publ 3799], fig. 2

iTtTbiriSoTroTroSiD
220131291--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Dai et al., 2023 [Publ 4285], fig. 3-ML

iTtTbiriSoTroTroSiD
220243431--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Lopes-Lima et al., 2020 [Publ 4080], fig. 9

iTtTbiriSoTroTroSiD
2201441--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Wu et al., 2024 [Publ 4388], fig. 6

iTtTbiriSoTroTroSiD
220152521--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Dai et al., 2025 [Publ 4440], fig. 1

iTtTbiriSoTroTroSiD
220243431--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Bespalaya et al., 2018 [Publ 3811], fig. 3

iTtTbiriSoTroTroSiD
2202881--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Lopes-Lima et al., 2019 [Publ 3901], fig. 2

iTtTbiriSoTroTroSiD
220126261--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Pfeiffer & Graf, 2015 [Publ 3620], fig. 2

iTtTbiriSoTroTroSiD
220138301--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lopes-Lima et al., 2020 [Publ 4080], fig. 16

iTtTbiriSoTroTroSiD
2201661--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Hou et al., 2024 [Publ 4400], fig. 4

iTtTbiriSoTroTroSiD
220174741--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lopes-Lima et al., 2020 [Publ 4080], fig. 10

iTtTbiriSoTroTroSiD
220112121--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Hoeh et al., 2009 [Publ 2716], fig. 5

iTtTbiriSoTroTroSiD
220144241--
1

Guerra et al., 2017 [Publ 3772], fig. 2b

iTtTbiriSoTroTroSiD
220123191--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Dai et al., 2024 [Publ 4385], fig. 3-ML

iTtTbiriSoTroTroSiD
220229291--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Välilä et al., 2017 [Publ 3793], fig. 2

iTtTbiriSoTroTroSiD
2201111--
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Hoeh et al., 2002 [Publ 2653], fig. 2

iTtTbiriSoTroTroSiD
220129181--
  +----------- Parreysiinae    
--+                            
  |  +-------- Margaritiferinae **
  +--+                         
     |  +----- Ambleminae      
     +--+                      
        |  +-- Gonideinae      
        +--+                   
           +-- Unioninae

Chen et al., 2023 [Publ 4302], fig. 1

iTtTbiriSoTroTroSiD
220238381--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wu et al., 2024 [Publ 4379], fig. 4_6

iTtTbiriSoTroTroSiD
220120201--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Dai et al., 2025 [Publ 4440], fig. 2

iTtTbiriSoTroTroSiD
220243431--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wu et al., 2024 [Publ 4350], fig. 6-7

iTtTbiriSoTroTroSiD
220135351--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Huang et al., 2019 [Publ 3843], fig. S1

iTtTbiriSoTroTroSiD
220243431--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wu et al., 2024 [Publ 4388], fig. 5

iTtTbiriSoTroTroSiD
220136361--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Krebs et al., 2003 [Publ 2691], fig. 3

iTtTbiriSoTroTroSiD
220131311--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Graf & Cummings, 2006 [Publ 1968], fig. 2

iTtTbiriSoTroTroSiD
220145251--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Pfeiffer et al., 2019 [Publ 3937], fig. 6

iTtTbiriSoTroTroSiD
220155501--
1

Lopes-Lima et al., 2020 [Publ 4080], fig. 12

iTtTbiriSoTroTroSiD
2201661--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lopes-Lima et al., 2020 [Publ 4080], fig. 6

iTtTbiriSoTroTroSiD
2201991--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Riccardi et al., 2020 [Publ 4089], fig. 2

iTtTbiriSoTroTroSiD
220117171--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Wu et al., 2023 [Publ 4339], fig. 5

iTtTbiriSoTroTroSiD
220117171--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Amaro et al., 2017 [Publ 3791], fig. 3B

iTtTbiriSoTroTroSiD
22011091--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Wu et al., 2018 [Publ 3836], fig. 2

iTtTbiriSoTroTroSiD
220151511--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Wu et al., 2024 [Publ 4350], fig. 8b_S2

iTtTbiriSoTroTroSiD
220189851--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lopes-Lima et al., 2019 [Publ 3901], fig. 3

iTtTbiriSoTroTroSiD
220127271--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Dai et al., 2023 [Publ 4285], fig. 3-BI

iTtTbiriSoTroTroSiD
220243431--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Chen et al., 2023 [Publ 4302], fig. 2

iTtTbiriSoTroTroSiD
220238381--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Huang et al., 2019 [Publ 3843], fig. 3-M

iTtTbiriSoTroTroSiD
220121201--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Dai et al., 2024 [Publ 4358], fig. 4

iTtTbiriSoTroTroSiD
220250501--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Hoeh et al., 2009 [Publ 2716], fig. 4

iTtTbiriSoTroTroSiD
220144241--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Wu et al., 2024 [Publ 4350], fig. 5b

iTtTbiriSoTroTroSiD
220135351--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Graf & Ó Foighil, 2000 [Publ 2648], fig. 2B

iTtTbiriSoTroTroSiD
220128251--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Konopleva et al., 2019 [Publ 3900], fig. 1

iTtTbiriSoTroTroSiD
220250501--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Lopes-Lima et al., 2020 [Publ 4080], fig. 17

iTtTbiriSoTroTroSiD
2201441--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Wu et al., 2024 [Publ 4350], fig. 5a

iTtTbiriSoTroTroSiD
220135351--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Amaro et al., 2017 [Publ 3791], fig. 2

iTtTbiriSoTroTroSiD
215011413133
  +-------- Margaritifera_margaritifera__2 **
--+                                       
  |  +----- Unionidae                     
  +--+                                    
     |  +-- Margaritifera_margaritifera__1 **
     +--+                                 
        +-- Margaritifera_falcata **

Campbell et al., 2005 [Publ 2660], fig. 1

iTtTbiriSoTroTroSiD
22011001001--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lopes-Lima et al., 2020 [Publ 4080], fig. 5

iTtTbiriSoTroTroSiD
2201771--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Dai et al., 2024 [Publ 4385], fig. 3-BI

iTtTbiriSoTroTroSiD
220229291--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Roe & Hoeh, 2003 [Publ 2655], fig. 4f3

iTtTbiriSoTroTroSiD
220129181--
1

Whelan et al., 2011 [Publ 3380], fig. 1-MP

iTtTbiriSoTroTroSiD
220134241--
1

Bogan et al., 2023 [Publ 4295], fig. 3

iTtTbiriSoTroTroSiD
220229291--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Graf & Ó Foighil, 2000 [Publ 2648], fig. 2A

iTtTbiriSoTroTroSiD
220128251--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Zieritz et al., 2021 [Publ 4218], fig. 5

iTtTbiriSoTroTroSiD
220130301--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Whelan et al., 2011 [Publ 3380], fig. 1-ML

iTtTbiriSoTroTroSiD
220134241--
1

Huang et al., 2019 [Publ 3843], fig. 1

iTtTbiriSoTroTroSiD
220241411--
  +----- Unionidae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Gibbosulinae **

Lopes-Lima et al., 2020 [Publ 4080], fig. 7

iTtTbiriSoTroTroSiD
2201661--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Fonseca et al., 2016 [Publ 3775], fig. 1

iTtTbiriSoTroTroSiD
220129291--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Teiga-Teixeira et al., 2020 [Publ 4173], fig. 1

iTtTbiriSoTroTroSiD
220117171--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Bogan & Hoeh, 2000 [Publ 2650], fig. 1

iTtTbiriSoTroTroSiD
220132181--
  +-------- Anodontini      
  |                         
--+  +----- Gonideinae      
  |  |                      
  +--+----- Unionini        
     |                      
     |  +-- Margaritiferinae **
     +--+                   
        +-- Ambleminae

Wen et al., 2017 [Publ 3795], fig. 3

iTtTbiriSoTroTroSiD
220114141--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Vikhrev et al., 2017 [Publ 3810], fig. 4

iTtTbiriSoTroTroSiD
2202881--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Lopes-Lima et al., 2020 [Publ 4080], fig. 11

iTtTbiriSoTroTroSiD
220110101--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lopes-Lima et al., 2020 [Publ 4080], fig. 13

iTtTbiriSoTroTroSiD
2201881--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Liu et al., 2024 [Publ 4380], fig. 5

iTtTbiriSoTroTroSiD
220135351--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Pfeiffer et al., 2019 [Publ 3937], fig. 4-5

iTtTbiriSoTroTroSiD
220155501--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lopes-Lima et al., 2020 [Publ 4080], fig. 2-4

iTtTbiriSoTroTroSiD
22011191101--
1

Bolotov et al., 2016 [Publ 3732], fig. 2

iTtTbiriSoTroTroSiD
2202661--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Feng et al., 2016 [Publ 3800], fig. 2

iTtTbiriSoTroTroSiD
220112121--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Campbell et al., 2005 [Publ 2660], fig. 2

iTtTbiriSoTroTroSiD
22011001001--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Dai et al., 2025 [Publ 4441], fig. 1

iTtTbiriSoTroTroSiD
220246461--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Wu et al., 2024 [Publ 4350], fig. 8a_S1

iTtTbiriSoTroTroSiD
220189851--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Zhang et al., 2024 [Publ 4397], fig. 5

iTtTbiriSoTroTroSiD
220126261--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lopes-Lima et al., 2020 [Publ 4080], fig. 14

iTtTbiriSoTroTroSiD
2201441--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Machordom et al., 2003 [Publ 2688], fig. 2

iTtTbiriSoTroTroSiD
2201111--
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Lopes-Lima et al., 2017 [Publ 3734], fig. 1

iTtTbiriSoTroTroSiD
220115141--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Wu et al., 2024 [Publ 4350], fig. 5c

iTtTbiriSoTroTroSiD
220135351--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lopes-Lima et al., 2017 [Publ 3745], fig. 1-2

iTtTbiriSoTroTroSiD
220178681--
1

Guerra et al., 2017 [Publ 3772], fig. 2a

iTtTbiriSoTroTroSiD
220123191--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Pavan-Kumar et al., 2022 [Publ 4263], fig. S5

iTtTbiriSoTroTroSiD
220136361--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lopes-Lima et al., 2019 [Publ 3901], fig. 4

iTtTbiriSoTroTroSiD
220124241--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Huang et al., 2002 [Publ 2657], fig. 1

iTtTbiriSoTroTroSiD
234013232140
  +-- Cumberlandia ** 
  |                
--+-- Margaritifera **
  |                
  +-- Unionidae

Dai et al., 2024 [Publ 4386], fig. 5-BI

iTtTbiriSoTroTroSiD
220219191--
  +----- Unionidae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Gibbosulinae **

Dai et al., 2024 [Publ 4386], fig. 5-ML

iTtTbiriSoTroTroSiD
220219191--
  +----- Unionidae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Gibbosulinae **

Sano et al., 2017 [Publ 3799], fig. 1

iTtTbiriSoTroTroSiD
220119171--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Hoeh et al., 2001 [Publ 2652], fig. 14f2

iTtTbiriSoTroTroSiD
220129181--
1

Burzynski & Soroka, 2018 [Publ 3862], fig. 5a

iTtTbiriSoTroTroSiD
220118181--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Bolotov et al., 2022 [Publ 4250], fig. 8

iTtTbiriSoTroTroSiD
27024747125
     +----- Parreysiinae    
  +--+                      
  |  |  +-- Pseudodontini   
  |  +--+                   
  |     +-- Contradentini   
  |                         
--+     +-- Gibbosulinae **    
  |     |                   
  |     +-- Margaritiferinae **
  |     |                   
  |     +-- Sinosolenaia    
  |     |                   
  +-----+-- Leguminaia      
        |                   
        +-- Gonidea         
        |                   
        +-- Lamprotula      
        |                   
        +-- Potomida

Roe & Hoeh, 2003 [Publ 2655], fig. 4f2

iTtTbiriSoTroTroSiD
220129181--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Dai et al., 2024 [Publ 4389], fig. 3

iTtTbiriSoTroTroSiD
220245451--
  +----- Unionidae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Gibbosulinae **

Hoeh et al., 2009 [Publ 2716], fig. 3

iTtTbiriSoTroTroSiD
220145251--
1

Pavan-Kumar et al., 2022 [Publ 4263], fig. 4

iTtTbiriSoTroTroSiD
220134341--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Dai et al., 2023 [Publ 4285], fig. 5

iTtTbiriSoTroTroSiD
220251511--
  +----- Unionidae       
--+                      
  |  +-- Gibbosulinae **    
  +--+                   
     +-- Margaritiferinae **

Amaro et al., 2017 [Publ 3791], fig. 3A

iTtTbiriSoTroTroSiD
22011091--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Araujo et al., 2009 [Publ 2589], fig. 7B

iTtTbiriSoTroTroSiD
110110101--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Pfeiffer & Graf, 2013 [Publ 3531], fig. 2

iTtTbiriSoTroTroSiD
110130291--
-- Margaritifera_margaritifera **

Marshall et al., 2014 [Publ 3593], fig. 1

iTtTbiriSoTroTroSiD
11011331--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Combosch et al., 2017 [Publ 3806], fig. 1

iTtTbiriSoTroTroSiD
11011431--
-- Margaritifera_margaritifera **

Chase et al., 2018 [Publ 3861], fig. 1

iTtTbiriSoTroTroSiD
11011091--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Hewitt et al., 2018 [Publ 3758], fig. 5

iTtTbiriSoTroTroSiD
1101881--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Hoeh et al., 1998 [Publ 2695], fig. 3

iTtTbiriSoTroTroSiD
1101421--
  +----- Unioninae       
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Ambleminae

Froufe et al., 2019 [Publ 3936], fig. 4

iTtTbiriSoTroTroSiD
110118181--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Curole & Kocher, 2005 [Publ 2682], fig. 1-M

iTtTbiriSoTroTroSiD
110111111--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Pfeiffer et al., 2018 [Publ 3827], fig. 2

iTtTbiriSoTroTroSiD
110127271--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Froufe et al., 2019 [Publ 3936], fig. 3-M

iTtTbiriSoTroTroSiD
110118181--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Inoue et al., 2018 [Publ 3858], fig. 1_3a-c

iTtTbiriSoTroTroSiD
11011031031--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Guerra et al., 2017 [Publ 3772], fig. 2b-M

iTtTbiriSoTroTroSiD
1101761--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Zanatta & Murphy, 2006 [Publ 2672], fig. 1

iTtTbiriSoTroTroSiD
110155551--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lydeard et al., 2000 [Publ 2654], fig. 2c

iTtTbiriSoTroTroSiD
110113131--
  +----- Unioninae       
  |                      
  +----- Margaritiferinae **
  |                      
--+----- Quadrulini      
  |                      
  +----- Lampsilini      
  |                      
  |  +-- Amblemini       
  +--+                   
     +-- Pleurobemini

Graf & Ó Foighil, 2000 [Publ 2649], fig. 1_2

iTtTbiriSoTroTroSiD
11011041--
-- Cumberlandia_monodonta **

Källersjö et al., 2005 [Publ 2684], fig. 1A

iTtTbiriSoTroTroSiD
1101661--
  +-- Margaritiferinae **
  |                   
--+-- Anodontini      
  |                   
  +-- Unionini

Wu et al., 2020 [Publ 4147], fig. 3

iTtTbiriSoTroTroSiD
110128281--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Prié & Puillandre, 2014 [Publ 3529], fig. 2

iTtTbiriSoTroTroSiD
110116161--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Smith, 2021 [Publ 4331], fig. 1

iTtTbiriSoTroTroSiD
110119161--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lemer et al., 2019 [Publ 4093], fig. 1

iTtTbiriSoTroTroSiD
1101511--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Reis et al., 2013 [Publ 3580], fig. 5

iTtTbiriSoTroTroSiD
110114131--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Chen et al., 2020 [Publ 4170], fig. 1

iTtTbiriSoTroTroSiD
110133331--
                       +----- Prolasmidonta           
                    +--+                              
                    |  |  +-- Alasmidonta             
                    |  +--+                           
  +-----------------+     +-- Anemina                 
  |                 |                                 
  |                 |  +----- Pseudodontini           
  |                 +--+                              
  |                    |  +-- Lamprotula_caveata      
  |                    +--+                           
  |                       +-- Chamberlainia           
  |                                                   
  |                    +----- Schistodesmus           
  |              +-----+                              
  |              |     |  +-- Lepidodesmini           
--+              |     +--+                           
  |              |        +-- Aculamprotula           
  |  +-----------+                                    
  |  |           |     +----- Platynaias              
  |  |           |  +--+                              
  |  |           |  |  |  +-- Utterbackia_imbecillis  
  |  |           |  |  +--+                           
  |  |           +--+     +-- Pyganodon               
  |  |              |                                 
  |  |              |  +----- Sinanodonta             
  |  |              +--+                              
  |  |                 |  +-- Cristaria               
  +--+                 +--+                           
     |                    +-- Anodontina              
     |                                                
     |                    +-- Utterbackia_peninsularis
     |              +-----+                           
     |              |     +-- Potamilus_alatus        
     |  +-----------+                                 
     |  |           |  +----- Lamprotula_leaii        
     |  |           +--+                              
     |  |              |  +-- Sinohyriopsis           
     +--+              +--+                           
        |                 +-- Gonideini               
        |                                             
        |  +----------------- Gibbosulinae **            
        +--+                                          
           |  +-------------- Quadrulini              
           +--+                                       
              |  +----------- Toxolasma               
              +--+                                    
                 |  +-------- Lampsilis_ornata        
                 +--+                                 
                    |  +----- Potamilus_leptodon      
                    +--+                              
                       |  +-- Lampsilis_siliquoidea   
                       +--+                           
                          +-- Lampsilis_powellii

Combosch et al., 2017 [Publ 3806], fig. 2

iTtTbiriSoTroTroSiD
11011431--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Walker et al., 2006 [Publ 2666], fig. 2

iTtTbiriSoTroTroSiD
110114121--
  +-------------- Unioninae       
  |                               
--+  +----------- Lamprotulini    
  |  |                            
  +--+        +-- Gonideini       
     |  +-----+                   
     |  |     +-- Pseudodontini   
     +--+                         
        |  +----- Lampsilini      
        +--+                      
           |  +-- Pleurobemini    
           +--+                   
              +-- Margaritiferinae **

Konopleva et al., 2023 [Publ 4269], fig. S1_4

iTtTbiriSoTroTroSiD
110155551--
  +-- Gibbosulinae **
--+               
  +-- Unionidae

Graf, 2000 [Publ 2647], fig. 1

iTtTbiriSoTroTroSiD
11011661--
-- Margaritifera **

Bogatov et al., 2018 [Publ 3859], fig. 1

iTtTbiriSoTroTroSiD
1101551--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Zanatta & Murphy, 2006 [Publ 2672], fig. 2

iTtTbiriSoTroTroSiD
110156561--
  +-- Unioninae       
  |                   
--+-- Margaritiferinae **
  |                   
  +-- Ambleminae

Graf et al., 2015 [Publ 3619], fig. 1

iTtTbiriSoTroTroSiD
11011911--
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Bogan et al., 2023 [Publ 4295], fig. 6

iTtTbiriSoTroTroSiD
110112121--
  +-- Gibbosulinae **
--+               
  +-- Unionidae

González & Giribet, 2015 [Publ 3632], fig. 4

iTtTbiriSoTroTroSiD
1101511--
  +-- Unionidae       
--+                   
  +-- Margaritiferinae **

Huang et al., 2018 [Publ 3828], fig. 6-M

iTtTbiriSoTroTroSiD
110116151--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Campbell, 2000 [Publ 2724], fig. 1-5

iTtTbiriSoTroTroSiD
1101221--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Curole & Kocher, 2002 [Publ 2712], fig. 1-F

iTtTbiriSoTroTroSiD
1101331--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lopes-Lima et al., 2017 [Publ 3792], fig. 2-M

iTtTbiriSoTroTroSiD
1101991--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Inoue et al., 2014 [Publ 3635], fig. 3A

iTtTbiriSoTroTroSiD
110119191--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Araujo et al., 2009 [Publ 2589], fig. 7C

iTtTbiriSoTroTroSiD
1101991--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Graf, 2002 [Publ 2651], fig. 2

iTtTbiriSoTroTroSiD
110117121--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Zieritz et al., 2012 [Publ 3673], fig. 4

iTtTbiriSoTroTroSiD
1101771--
  +----- Anodontina      
  |                      
--+  +-- Cristariina     
  |  |                   
  +--+-- Margaritiferinae **
     |                   
     +-- Unionini

Chase et al., 2018 [Publ 3861], fig. 1-M

iTtTbiriSoTroTroSiD
11011091--
-- Cumberlandia_monodonta **

Bogatov et al., 2018 [Publ 3859], fig. 2

iTtTbiriSoTroTroSiD
1101551--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Matos et al., 2024 [Publ 4384], fig. 3

iTtTbiriSoTroTroSiD
110119191--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Huang et al., 2013 [Publ 3629], fig. 3

iTtTbiriSoTroTroSiD
110115151--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Zieritz et al., 2021 [Publ 4231], fig. 3-M_4

iTtTbiriSoTroTroSiD
110122221--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lydeard et al., 2000 [Publ 2654], fig. 2a

iTtTbiriSoTroTroSiD
110113131--
  +----- Unioninae       
  |                      
  +----- Margaritiferinae **
  |                      
--+----- Lampsilini      
  |                      
  +----- Amblemini       
  |                      
  |  +-- Quadrulini      
  +--+                   
     +-- Pleurobemini

Hoeh et al., 1998 [Publ 2695], fig. 4

iTtTbiriSoTroTroSiD
1101421--
  +----- Ambleminae      
--+                      
  |  +-- Margaritiferinae **
  +--+                   
     +-- Unioninae

Taylor et al., 2007 [Publ 2696], fig. 1-3

iTtTbiriSoTroTroSiD
1101321--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Bogan et al., 2023 [Publ 4295], fig. 5

iTtTbiriSoTroTroSiD
1101551--
  +-- Gibbosulinae **
--+               
  +-- Unionidae

Bieler et al., 2014 [Publ 3622], fig. 28

iTtTbiriSoTroTroSiD
1101411--
-- Margaritifera_margaritifera **

Graf et al., 2014 [Publ 3591], fig. 3

iTtTbiriSoTroTroSiD
110110101--
  +-- Margaritiferinae **
  |                   
--+-- Unioninae       
  |                   
  +-- Parreysiinae

Froufe et al., 2017 [Publ 3802], fig. 7

iTtTbiriSoTroTroSiD
1101991--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Walker et al., 2006 [Publ 2666], fig. 3

iTtTbiriSoTroTroSiD
110114121--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Araujo et al., 2018 [Publ 3779], fig. 1

iTtTbiriSoTroTroSiD
110117171--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Amaro et al., 2017 [Publ 3791], fig. 3A-M

iTtTbiriSoTroTroSiD
1101661--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Douda et al., 2024 [Publ 4364], fig. 1

iTtTbiriSoTroTroSiD
110113131--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Nagel et al., 1998 [Publ 1946], fig. 9

iTtTbiriSoTroTroSiD
1101551--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Källersjö et al., 2005 [Publ 2684], fig. 1B-F

iTtTbiriSoTroTroSiD
1101661--
  +-- Margaritiferinae **
  |                   
--+-- Anodontini      
  |                   
  +-- Unionini

Gustafson & Iwamoto, 2005 [Publ 2741], fig. 2

iTtTbiriSoTroTroSiD
1101441--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Lydeard et al., 2000 [Publ 2654], fig. 2d

iTtTbiriSoTroTroSiD
110113131--
  +-- Unioninae       
  |                   
--+-- Margaritiferinae **
  |                   
  +-- Ambleminae

Curole & Kocher, 2005 [Publ 2682], fig. 1

iTtTbiriSoTroTroSiD
110110101--
  +----- Ambleminae      
--+                      
  |  +-- Unioninae       
  +--+                   
     +-- Margaritiferinae **

Reis et al., 2013 [Publ 3580], fig. 4

iTtTbiriSoTroTroSiD
110113121--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Prie et al., 2012 [Publ 3487], fig. 2

iTtTbiriSoTroTroSiD
1101551--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

González & Giribet, 2015 [Publ 3632], fig. 2

iTtTbiriSoTroTroSiD
1101511--
-- Margaritifera_margaritifera **

Bogan et al., 2023 [Publ 4295], fig. 4

iTtTbiriSoTroTroSiD
1101661--
  +-- Gibbosulinae **
--+               
  +-- Unionidae

Zhang et al., 2013 [Publ 4214], fig. 5

iTtTbiriSoTroTroSiD
1101331--
  +-------- Cristaria       
--+                         
  |  +----- Sinanodonta     
  +--+                      
     |  +-- Margaritiferinae **
     +--+                   
        +-- Gonideinae

Hewitt et al., 2019 [Publ 3962], fig. 2

iTtTbiriSoTroTroSiD
1101991--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Araujo et al., 2017 [Publ 3801], fig. 2

iTtTbiriSoTroTroSiD
110113131--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Matos et al., 2024 [Publ 4383], fig. 3

iTtTbiriSoTroTroSiD
110119191--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Klishko et al., 2019 [Publ 3886], fig. 9

iTtTbiriSoTroTroSiD
110110101--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Kano et al., 2019 [Publ 4194], fig. 6

iTtTbiriSoTroTroSiD
1101991--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Walker et al., 2006 [Publ 2666], fig. 4

iTtTbiriSoTroTroSiD
110114121--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Amaro et al., 2017 [Publ 3791], fig. 3B-M

iTtTbiriSoTroTroSiD
1101661--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Guerra et al., 2017 [Publ 3772], fig. 2a-M

iTtTbiriSoTroTroSiD
1101761--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Turner et al., 2000 [Publ 2722], fig. 2

iTtTbiriSoTroTroSiD
110113131--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Araujo et al., 2009 [Publ 2589], fig. 7A

iTtTbiriSoTroTroSiD
110110101--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

González & Giribet, 2015 [Publ 3632], fig. 3

iTtTbiriSoTroTroSiD
1101511--
-- Margaritifera_margaritifera **

Lydeard et al., 2000 [Publ 2654], fig. 2b

iTtTbiriSoTroTroSiD
110113131--
  +-- Unioninae       
  |                   
--+-- Margaritiferinae **
  |                   
  +-- Ambleminae

González & Giribet, 2015 [Publ 3632], fig. 6

iTtTbiriSoTroTroSiD
1101511--
-- Margaritifera_margaritifera **

Bieler et al., 2014 [Publ 3622], fig. 26_27_29_30

iTtTbiriSoTroTroSiD
1101411--
-- Margaritifera_margaritifera **

Chase et al., 2018 [Publ 3847], fig. 5

iTtTbiriSoTroTroSiD
1101981--
-- Cumberlandia_monodonta **

Klunzinger et al., 2021 [Publ 4086], fig. 2

iTtTbiriSoTroTroSiD
1101511--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Bashê, 2022 [Publ 4265], fig. 1

iTtTbiriSoTroTroSiD
1101551--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Giribet & Distel, 2003 [Publ 3670], fig. 3f5

iTtTbiriSoTroTroSiD
1101531--
  +-- Margaritiferinae **
--+                   
  +-- Unionidae

Zhou et al., 2016 [Publ 3710], fig. 1

iTtTbiriSoTroTroSiD
110119191--
  +-------- Unioninae       
--+                         
  |  +----- Margaritiferinae **
  +--+                      
     |  +-- Ambleminae      
     +--+                   
        +-- Gonideinae
 
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